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A New Genus from Antarctic Waters, with the Description of a New Species

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A New Genus from Antarctic Waters, with the Description of a New Species View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by CONICET Digital THE VELIGER # The Veliger 51(1):85–103 (January/April XX, 2008) CMS, Inc., 2008 Trophonella (Gastropoda: Muricidae), a New Genus from Antarctic Waters, with the Description of a New Species M. G. HARASEWYCH Department of Zoology, MRC-163, National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, Washington, DC 20013-7012, USA (e-mail: [email protected]) GUIDO PASTORINO Museo Argentino de Ciencias Natureles, Av. Angel Gallardo 470, 3u piso, lab. 57, C1405DJR Buenos Aires, Argentina (e-mail: [email protected]) Abstract. The new genus Trophonella is described from the outer shelf and upper continental slope of Antarctica and islands within the Antarctic Convergence. Four previously known species that had been attributed to the genus Trophon (Trophon scotianus Powell, 1951; T. echinolamellatus Powell, 1951; T. enderbyensis Powell, 1958; and T. eversoni Houart, 1997) are included in Trophonella, as is one new species (Trophonella rugosolamellata) described herein. Trophonella resembles Trophon in gross shell morphology: the members of both genera have large, globose shells, paucispiral protoconchs, prominent axial lamellae, and short siphonal canals. Trophonella differs from Trophon in having shells with evenly rounded whorls that lack a well-defined shoulder; rachidian teeth with distinctive, broadly triangular central cusps, but that lack the marginal cusps of Trophon; characteristic spherical accessory salivary glands; and a circumpapillar fold on the penis that is absent in Trophon. Relationships of the genera Trophon and Trophonella, as well as of the subfamily Trophoninae are reexamined by supplementing the data matrix of Kool (1993b, Table 3) with data for additional taxa. Results support the segregation of Trophonella from Trophon at the generic level. Based on the relationships of the type species of their respective nominotypical genera, Trophoninae is either the sister taxon of a narrowly circumscribed Ocenebrinae, or both are part of a larger clade. A better resolved phylogeny containing a much broader sampling of the more than 50 genus-level taxa that have been attributed to these two subfamilies will be required in order to delineate more precisely the membership of the clade and to identify its diagnostic synapomorphies. INTRODUCTION site of assignment of many dissimilar forms that seem, on the basis of the shell and radular morphology of the Although the family Muricidae is best known for its type species of Trophon (T. geversianus), unlikely to temperate and tropical shallow-water representatives, it belong there.’’ The genus Aspella is presently assigned also includes a group of taxa inhabiting polar seas and to the subfamily Muricinae on the basis of radular intervening deeper waters that have generally been morphology (Radwin & D’Attilio, 1976, p. 21). These grouped in the subfamily Trophoninae. Cossmann authors went on to exclude from the subfamily (1903) proposed the subfamily Trophoninae to contain Trophoninae a number of genera (e.g., Austrotrophon the genera Trophon Montfort, 1810, and Aspella Dall, 1902; Forreria; Zacatrophon Hertlein & Strong, Mo¨rch, 1877. The genus Trophon was subdivided into 1951) because they ‘‘… had radulae characteristic of the the subgenera Trophon and Trophonopsis Bucquoy and family Thaididae.’’ In a phylogenetic study, Kool Dautzenberg, 1882. Under Trophon sensu stricto,he (1993a, fig. 65) showed that, based on the morphology included the sections Trophon, Xanthochorus Fischer, of its type species, the genus Trophon was closely 1884, and Forreria Jouseaume, 1880. Trophonopsis was related to the genera Nucella and Ocenebra,andhe further subdivided into Trophonopsis and Boreotrophon suggested that future studies would reveal Trophoninae Fischer, 1884. The genus Aspella was not subdivided. to be polyphyletic. Over the past several decades, there has been general Our continuing studies on the Muricidae represented agreement among researchers that Trophoninae is not in the collections of the United States Antarctic a monophyletic group. As was noted by Radwin and Program (USAP) have revealed that four species D’Attilio (1976, p. 175), Trophoninae ‘‘has been the previously described in the genus Trophon (i.e., The Veliger veli-51-01-08.3d 24/11/09 04:21:36 85 Page 86 The Veliger, Vol. 51, No. 1 Trophon scotianus Powell, 1951; T. echinolamellatus phon geversianus, Ocenebra erinaceus, Trophonella Powell, 1951; T. enderbyensis Powell, 1958; and T. scotiana, Boreotrophon aculeatus,andPaziella pazi. eversoni Houart, 1997) appear to be closely related to Harasewych (1984) was the source of data for T. each other, but they differ substantially from the type geversianus, B. aculeatus,andP. pazi.DataforT. species of the genus Trophon. In this paper, we propose geversianus and O. erinaceus were obtained from Kool the new genus Trophonella to include these species, and (1993a). Data on shell ultrastructure of Boreotrophon we describe an additional new species that we attribute aculeatus and Paziella pazi were newly obtained from to this genus. A review of each of the species included the voucher material from Harasewych’s (1984) study. in this new genus is provided, based on more recently Each of the newly added taxa were scored for the 18 collected specimens. characters and their character states, as defined in Kool We also examine the relationships of the genus (1993b). The data were analyzed using maximum Trophon and the subfamily Trophoninae by reanalyz- parsimony (MP) and neighbor-joining algorithms of ing Kool’s (1993b, table 3) data matrix of primarily PAUP* 4.0 Macintosh Beta Version 10 (Swofford, ocenebrine and rapanine taxa supplemented by the 2002). addition of data for (1) Trophon geversianus (Pallas, 1769), the type species of the type genus of Trophoni- SYSTEMATICS nae; (2) Ocenebra erinaceus (Linnaeus, 1758), the type species of the type genus of Ocenebrinae; (3) Tropho- Class GASTROPODA Cuvier, 1797 nella scotiana (Powell, 1951), the type species of the new Order NEOGASTROPODA Wenz, 1938 genus Trophonella; (4) Boreotrophon aculeatus Watson, 1882; and (5) Paziella pazi (Crosse, 1869), the type Family MURICIDAE Rafinesque, 1815 species of the genus Paziella. Trophonella new genus MATERIALS AND METHODS Type species: Trophon scotianus Powell, 1951. This study is based primarily on specimens collected for Diagnosis: Shell large, fusiform, inflated. Protoconch the United States Antarctic Program (USAP) aboard paucispiral (1.5–2 whorls). Teleoconch (6–7 whorls) the vessels R/V Hero, R/V Eltanin and R/V Professor globose, without distinct shoulder, with tall, conical Sedlacki and housed in the collections of the National spire. Sculpture of low, rounded, spiral cords and axial Museum of Natural History, Smithsonian Institution lamellae that may be broadly flaring to short and (USNM). Additional specimens from several Antarctic rugose. Aperture broadly rounded, with flaring outer expeditions of the Argentine Republic deposited in the lip, and thin inner lip. Siphonal canal short. Operculum Museo Argentino de Ciencias Naturales (MACN) were large, oval, with subterminal nucleus. Rachidian teeth examined, as were specimens at the Zoological Institute with rectangular basal plate; three major cusps, central and Museum, Hamburg (ZMH), and the Senckenberg cusp exceptionally broad, triangular, with small denti- Museum, Frankfurt (SMF), collected by the vessels R/ cle on each side; marginal cusps absent. Accessory V Walther Herwig and R/V Polarstern. All specimens salivary glands small, spherical. Esophageal gland were compared with the holotypes of their respective reduced. Penis with conical papilla surrounded by species, which are housed in The Natural History collar. Museum, London (NHM), the South Australian Museum (SAM), and at USNM. Description: Shell large (to 76 mm), fusiform, often Alcohol-preserved animals of most species of Troph- broadly ovate, with strongly convex, evenly rounded onella were dissected, and the gross anatomy of the whorls that do not form a distinct shoulder. Proto- anterior part of the alimentary system and pallial conch paucispiral, occasionally with weak, irregular, gonoducts were compared. Radulae were prepared spiral cords. Teleoconch sculpture of low, rounded, according to the method described by Solem (1972) and spiral cords with or without scales, and axial lamellae examined using a Scanning Electron Microscope that range from broadly flaring to short and rugose. (SEM). Terminology for rachidian-tooth morphology Aperture large, suboval, with thick outer lip, and thin follows Kool (1987, fig. 1). Shell ultrastructure data inner lip. Siphonal canal short, scabrous. Shell was obtained from fracture surfaces of shell fragments ultrastructure of three layers, the inner two of removed from the central portion of the lip along the orthogonally oriented crossed-lamellar aragonite, the last shell whorl. outermost layer of calcite. Operculum large, filling In order to determine the generic and subfamilial aperture, corneous, oval, with subterminal nucleus, relationships of this distinctive group of Antarctic thick rim, and large attachment area. muricids, we reanalyzed the data matrix in Kool Animal large, unpigmented. Cephalic tentacles wide, (1993b, table 3), to which we added data for Tro- blunt. Mantle edge smooth, siphon short. Pleuroem- The Veliger veli-51-01-08.3d 24/11/09 04:21:36 86 M. G. Harasewych & G. Pastorino, 2008 Page
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